Abstract. Delimitation of genera in the Tovomita group of
tribe Clusieae has fluctuated, in Central America, with almost every
new floristic work. Nevertheless, with some aberrations, the theme of
a unified Chrysochlamys Tovomitopsis has persisted, and
is here supported by concurring with Bentham and Hooker's placement
of Tovomitopsis in synonymy of Chrysochlamys. The
strictly South American Balboa is also placed in synonymy of
Chrysochlamys, which action necessitates a new name (here
provided) for its only species. A key to the 13 species of
Chrysochlamys recognized for Mexico and Central America is
provided. An annotated list of all, including brief descriptions,
taxonomic and nomenclatural notes, five new combinations, and two new
species, brings forth that concept and rectifies the nomenclature to
the practicalities of numerous regional floras. Tovomita croatii Maguire, a somewhat problematic species, transferred
to Tovomitopsis by D'Arcy, is included in the key to
Chrysochlamys and fully discussed.

Taxonomic History

The Tovomita group of the tribe Clusieae (Clusiaceae,
subfamily Clusioideae) comprises primarily arborescent, terrestrial
genera with succulent-capsuled fruits and 1(rarely 2) large arillate
seeds per locule. Three genera therein, Tovomita,
Chrysochlamys, and Tovomitopsis have suffered a
particularly tortuous taxonomic history. Planchon and Triana (1860)
made the genus Tovomitopis to accommodate Sprengel's
Bertolonia, a later homonym; Bertolonia Raddi
(Melastomataceae) is now conserved over Bertolonia Spin
(Myoporaceae) (Farr et al., 1979). Planchon and Triana distinguished
Tovomitopsis from Chrysochlamys claiming it had an
arilloid (originating from the micropyle) rather than the true aril
(originating from the funicule) they claimed for
Chrysochlamys. According to them, the so-called aril of
Tovomita is neither, but rather the outer integument of the
seed. These differences were further supported by floral structure:
Tovomitopis, calyx 4-merous; Chrysochlamys, calyx
5-merous; Tovomita, calyx 2- or 4-parted, the outer two sepals
overtopping the bud. Planchon and Triana also described Balboa
in the same publication with Tovomitopsis, but they placed it
in their subtribe "Euclusiae" rather than Tovomiteae. Bentham and
Hooker (1862: 167--176) recognized Balboa,
Chrysochlamys and Tovomita (as related) but treated
Tovomitopsis as a synonym of Chrysochlamys. However,
they transferred no species. Subsequently, Balboa was
apparently forgotten in discussions relating to the Tovomita
group. Hemsley (1879: 87) transferred all Central American species of
Tovomitopsis to Chrysochlamys. Cuatrecasas (1950) also
declared unjustified the separation between Chrysochlamys and
Tovomitopsis and transferred two (South American) species to
Chrysochlamys. Williams (1959) subsumed Tovomitopsis
within Tovomita, but kept Chrysochlamys separate.
Maguire (1977) tentatively recognized all three genera. D'Arcy (1980)
expressed the same taxonomic opinion begun by Bentham and Hooker and
put into practice by Hemsley and Cuatrecasas, i.e., a unified
Chrysochlamys/Tovomitopsis, but used the later name,
Tovomitopsis. D'Arcy also elevated Engler's section
Dystovomita of Tovomita to the genus
Dystovomita.

Most recently, an attempt has been made to circumscribe
Chrysochlamys as comprising species with succulent petals,
echinate pollen exine, and black seeds with white arils, rather than
membranaceous petals, psilate-foveolate pollen, and green seeds with
orange arils (cf. Hammel, 1984, 1986). Ignoring whether or not these
characters support a monophyletic group or even if they coincide,
typologically, with Chrysochlamys, neither such a separation,
nor one based on the considerations of Planchon and Triana is
practical. Dystovomita, with axillary inflorescences (i.e.,
mostly lateral and below the leaves), large, adaxial petiolar pits
and relatively small seeds, is easily recognized and may even lie
closer to Clusia (P. Stevens, in prep.). Tovomita, with
terminal inflorescences, leaves clustered at the ends of side shoots,
leaves sometimes with petiolar pits, outer two sepals fused in bud or
at least overtopping the bud, styles often evident, fruits usually
with the inner wall and placenta dark reddish wine, can also be
recognized, even with most herbarium material. The older name,
Chrysochlamys, takes precedence over Tovomitopsis for
the remaining species in the group, i.e., those with terminal
inflorescences, leaves not clustered on side shoots, lacking adaxial
petiolar pits, sepals (usually) not overtopping the bud, styles
lacking, and the inner fruit wall and placenta mostly white or pink.
Balboa is a synonym of Chrysochlamys.

2a. Leaves clustered at the ends of side shoots; outer two
sepals fused in bud or at least overtopping the bud; styles often
evident; inner fruit wall and placenta usually dark reddish
wine......................................Tovomita

2b. Leaves ± evenly dispersed along the branches; outer
two sepals never fused in bud, nor both overtopping the bud; inner
fruit wall and placenta mostly white or
pink.........................................Chrysochlamys

Dioecious, terrestrial trees or shrubs, sometimes epipetric
rheophytes, the branching decussate and the leaves dispersed more or
less evenly along the branches; resin mostly clear, occasionally
white, rarely yellow; internodes mostly without bud-scale scars.
Petiolar pits and interpetiolar stipuliform structures absent. Leaves
opposite, the blades elliptic; major lateral veins mostly prominent,
resin canals sometimes visible in dried material. Inflorescences
terminal (rarely left pseudo-axillary by the dominant growth of an
axillary, vegetative bud), many-flowered, panicles. Flowers 4- or
5-merous, corolla aestivation imbricate, petals thin, membranaceous,
to thick and succulent, pink, or yellow to white. Staminate flowers
with numerous (rarely few) ± separate stamens, or the filaments
fused basally into a short column, or rarely with fertile stamens
surrounding a central sterile capitulum, or very rarely the anthers
all sessile on a resiniferous capitulum. Pistillate flowers with
numerous separate staminodia, or staminodia mounted on a ring or
rarely just a ring surrounding the ovary; ovary with mostly 4 or 5
stigmas, rarely 3. Fruits globose to ovoid, usually terete, rarely
winged or star-shaped in cross section, succulent capsules mostly
with 1 seed maturing per locule, sometimes only 1 seed per fruit;
seeds green to nearly black with an orange or white aril.

Distribution and ecology. Chrysochlamys is known
from Mexico to Bolivia and Brazil, where it occurs in wet to very wet
primary forest from near sea level to at least 2300 m.

Chrysochlamys, in total, contains perhaps as many as 50
species, most of them South American, and is in much need of
revision. At least two Central American species, C.
glauca (Oersted, Planchon & Triana) Hemsley and C.
nicaraguensis (Oersted, Planchon & Triana) Hemsley, have
been erroneously reported from South America, and numerous South
American names (e.g., C. membranacea Planchon &
Triana, C. micrantha Engler, C. myrcioides Planchon & Triana) have been loosely (and erroneously) applied to Central American species, as well as within South America. The following key, distinguishing 13 species of Chrysochlamys for Mexico and Central America, and the subsequent annotated list, including citation of representative specimens and descriptions of two new species from Costa Rica and Panama, aim to clarify the species and names for the region. One or two possible new species, from among Panamanian material at hand, await resolution and are not included here.

Key to the species of Chrysochlamys (and one
Tovomita) in Mesoamerica

1 Internodes with a pair of bud scales (or their scars),
usually just above the subtending node; primary lateral veins
close (ca. 3 mm apart), ± prominent but difficult to
distinguish from the intersecondaries; inflorescences often
appearing axillary; anthers sessile on a capitulum; stigmas 3; far
eastern Panama to
Ecuador...........................C.
tenuifolia

13' Fruits not alate; leaves oblong to
elliptic, not falcate, membranaceous, the apex
abruptly acute to acuminate, the resin canals
obscure; outer pair of sepals very unequal, the
smaller much shorter than the bud; resin canals of
petals obscure or lacking; resin of stem clear or
white.

Distribution. Very wet cloud forest, (700--)1450--2300 m;
continuous from the Cordillera de Guanacaste in northwestern Costa
Rica to Chiriquí province of western Panama, disjunct from
Darién province of far eastern Panama.

A larger tree than any other Chrysochlamys in Central
America and growing at higher elevation, C. allenii is
further distinguished by its coriaceous leaves, numerous stamens
(more than 50), often very large stigmas, and pyriform, often
greenish yellow fruits.

Although D'Arcy (1980) compared Tovomitopsis centistaminibus with T. psychotriifolia, the type specimen's high-elevation habitat, large stature, coriaceous leaves, and numerous stamens conspire to alienate it from the latter species, and instead, to bring it into synonymy with C. allenii. This species, however, can at times be confused with C. psychotriifolia, which usually grows at somewhat
lower elevations, has fewer (and often longer) stamens, and larger
fruits. Material at hand, especially that from far eastern Panama,
suggests that C. allenii may be closely related to,
perhaps not distinct from, C. colombiana (Cuatrecasas)
Cuatrecasas.

This distinctive, small, narrow-leaved, epipetric rheophyte with
flower buds over 6 mm long is endemic to a few lowland or mid-elevation sites in west central Panama. Its habitat, larger flowers, and stouter fruiting rachis easily distinguish it from the similarly narrow-leaved new species, C. tenuis, described below. Among Central American species, C. angustifolia is one of the few with copious milky resin rather than the more common clear to tardily creamy resin.

Distribution. Wet to very wet forest, from near sea level
to 1600 m; central to eastern Panama.

A distinctive, usually dark-reddish brown-drying, dull-leaved,
Panamanian endemic (from the Canal to San Blas and Darién)
with succulent and yellow rather than membranaceous white or pink
petals, stamens with short, free filaments mounted on a succulent,
fused basal portion, white rather than orange arils, and black rather
than green seeds, not at all closely related to C.
nicaraguensis, under which it was synonymized by D'Arcy
(1980). This is the only species, described from the region, with
succulent, rather than membranaceous petals. Even in fruit it can be
identified by the combination of large, dark-drying leaves and fused
staminodia. The petals dry black or very dark burgundy.

Some of this material with particularly large inflorescences from
far eastern Panama has been identified as C. floribunda
Cuatrecasas and C. weberbaueri Engler However, the
abundant material at hand suggests that probably only one species is
involved. Pending study of type material of the older, South American
names, I here use C. eclipes for all Panamanian
collections.

Distribution. Wet to very wet forest, from near sea level
to 1400 m; Costa Rica and Panama.

Even segregating the material described below as
Chrysochlamystenuis, C. glauca remains a
rather variable species, characterized by its usually dull,
few-veined leaves and small flower buds. Some of that variation may
be worthy of taxonomic recognition. Many of the collections from the
Osa peninsula in Costa Rica (cf. Aguilar431,
Kernan713) have leaves with especially obvious and
numerous latex canals and flower buds near the lower end of the size
range for the species, giving such material an aspect predictive of
its origin. Allen (1956: 345), in fact, distinguished two species
from the Osa region within the C. glauca complex. For
one (vouchered by Skutch5286, F, not seen), he used
the name C. costaricana, stating that it was "nearly
identical" to C. glauca, "differing principally in the
smaller flowers." Although C. costaricana (discussed
below under C. psychotriifolia) is not its correct
name, the species distinguished by Allen may deserve recognition, not
since given. Populations at the other end of Costa Rica, in the
Cordillera de Guanacaste, have almost shiny leaves with relatively
obscure lateral veins. The plants also appear to flower more on
lateral shoots than material from elsewhere. The anthers are slightly
longer and the stigmas are larger than those of C.
glauca from elsewhere, usually at least 1/2 (vs. no more than
1/3--1/4) the length of the ovary in newly opened flowers. Several
collections from Panama, Panama province and San Blas (deNevers7223, Herreraetal.
1137, Herreraetal. 1145),
tentatively placed here, have unusually dense and much-branched resin
lines, and fruits with persistent sepals. These may represent a
distinct species, but flowering material is needed.

Authorship of the four Central American Tovomitopsis names
published in Planchon and Triana (1860) has been variously obfuscated
in numerous publications, including my own. Oersted alone never
published these names, as sometimes implied (cf. Williams, 1959),
nor is there any basis for the even more common "Oersted ex Planchon
& Triana," (cf. Standley, 1937; Croat, 1978; D'Arcy, 1980), let
alone the terse permutation of the latter to just "Planchon &
Triana" (e.g., Hammel, 1986). Since Anders Örsted (1812--1872)
was professor of botany at the University of Copenhagen until 1862
(http://www.nathimus.ku.dk/bot/orsted.htm)
during the time Planchon and Triana were working in Paris on their
Guttiferae memoir, nothing suggests other than the three, "Oersted,
Planchon & Triana," as co-authors, as originally indicated for
all four species in Planchon and Triana (1860).

The citation of two collection (?) numbers for the types of these
same four species results from examination of the actual specimens.
The single-digit numbers are cited in the original publication (e.g.,
Oersted4), and are also written on or in packets with
the specimens. The four-digit numbers printed on the labels (e.g.,
Plantae Centroamericanae Oersted No. 3585) are not mentioned in the
publication. Neither series appears to come from normal consecutive
field-note numbers; earlier dates do not correspond, consistently, to
lower numbers within the series.

This distinctive Central American species, clearly delimited by
its protologue, is characterized by strongly ribbed fruits and
monomorphic stamens with anthers up to 2--3 times longer than wide.
Costa Rican material from the Pacific slope has glabrous
inflorescences and anthers nearly 2 mm long, whereas that of the
Caribbean slope (also occurring at higher elevation than that of the
Pacific) and the Panamanian collections have markedly puberulent
inflorescences and smaller anthers.

This species was originally published as Tovomitamacrophylla L. O. Williams, a homonym of the earlier T.
macrophylla (Poeppig) Walpers Williams, therefore, made a new
name, T. grandifolia, for the same species.
Coincidentally, an earlier Chrysochlamysmacrophylla
Pax would prevent the use of that combination for the present
species, in any case. Nothing, however, prevents the use of the
combination C. grandifolia (L. O. Williams) Hammel.

Although D'Arcy (1980) created the combination Tovomitopsismyrcioides (Planchon & Triana) D'Arcy for this material
(with Tovomitagrandifolia as synonym), the South
American C. myrcioides Planchon & Triana is a quite
different and poorly known species. It has very short petioles and
the dimorphic stamens of section Heterandra "externa fertilia,
libera, interna sterilia in corpus centrale concreta" (Planchon &
Triana, vol. 14: 260. 1860) with nearly globose anthers. Furthermore,
I know of no such rib-fruited Chrysochlamys species from South
America. Chrysochlamysmembrillensis (see below) of far
eastern Panama also has somewhat ribbed fruits, but is otherwise
quite different from the present species.

This name has been considered a synonym of Chrysochlamysnicaraguensis (cf. Standley & Williams, 1961) and
C. psychotriifolia (cf. Hammel, 1984). However, a
clearer picture of variation within the genus throughout the region
now supports its reinstatement. The species is characterized by more
or less shiny, usually reddish-brown-drying leaves with rather few
(6--9) and distant (1--2 cm) major lateral veins and usually quite
prominent intersecondaries and obvious resin lines. The rather large
flower buds (5--7 mm) have visible resin lines. It has more stamens
(ca. 50) than either C. nicaraguensis (35--40) or
C. psychotriifolia (ca. 20). The fruits are often quite
elongate, up to 4 cm, as in C. psychrotiifolia, and the
sepals are early deciduous.

Distribution. Very wet primary forest, 100--850 m. Known
for sure from just five collections, only from Panama, low to middle
elevations of Colón province and extreme eastern Darién
province. This species must certainly also occur in nearby Colombia,
but no such specimens have been seen.

This Panamanian endemic has narrowly elliptic and often falcate,
coriaceous leaves, with very visible resin canals. The lateral leaf
veins are without prominent intersecondaries, and the fruits are
basically pyriform and winged. Although the winged fruits are very
characteristic of this species, they are not obvious in young
pistillate material nor even more mature, dried material.

Distribution. Very wet forest of the Caribbean lowlands,
from near sea level to 600(--1200) m; Nicaragua, Costa Rica, Panama.

Chrysochlamys nicaraguensis, often erroneously
interpreted as a wide-ranging species (cf. discussion under C.
eclipes), is still only known for certain from the very wet
Caribbean lowlands of Nicaragua, Costa Rica, and adjacent Panama. It
is distinctive for its very glossy and distantly veined (2--3 cm)
leaves almost totally lacking visible intersecondaries and resin
canals. For discussion of authorship and type specimen citation of
this species, see under C. glauca.

Chrysochlamys psychotriifolia, well known in Costa
Rica, is a species with numerous, rather close, lateral veins, few
(ca. 20) monomorphic stamens, and usually large (3--5 cm),
obovate-mammillate fruits. Tovomitopsiscostaricana,
T. faucis, and what D'Arcy (1980) called T.
membranacea (Planchon & Triana) D'Arcy were based on very
scant material that is, nevertheless, now seen to fall easily within
the variation of C. psychotriifolia. For discussion of
authorship and type specimen citation of this species, see under
C. glauca. The South American Chrysochlamysmembranacea Planchon & Triana, by its type and numerous
more recent collections is clearly a different species, belonging to
Planchon and Triana's section Heterandra, with dimorphic
stamens in the staminate flowers and the staminodia connate into a
ring or collar in the pistillate.

This species, particularly as to its type, can be characterized by
its primary lateral veins and intersecondaries being almost equally
salient. Nevertheless, a few collections from Costa Rica included
here (Heredia, Cantón de Sarapiquí, Rara Avis, 400--700
m, Martén948 (CR); G. Vargas1650 (CR); O. Vargas2 (CR); O.
Vargas4 (CR)) have fewer and less prominent
intersecondaries. Material from Panama appears to grade imperceptibly
into an extreme with the intersecondaries very obscure. All the
material has dull, yellowish to reddish tan-drying leaves, the outer
pair of sepals often very unequal, the petals usually without obvious
resin canals, and stamens about 20--30. At least in Costa Rica, the
resin of the twigs is bright white, rather than clear or tardily
creamy as in most other species in Central America. For its reddish
tan-drying leaves, this species sometimes resembles the more northern
C. guatemaltecana, which has more stamens and shiny
leaves with fewer, more distant veins.

Distribution. This species is known only from a few
collections from relatively low 100--750(--900) m elevations in wet
forest on the Pacific slope of southern (and eastern) Costa Rica,
from the regions of Acosta (Fila Bustamante), Valle del General, the
Osa Peninsula and vicinity.

Flowering or sterile collections of this species may be difficult
to separate from Chrysochlamysgrandifolia, a species
most readily distinguished by its strongly ribbed fruits. The two
grow sypatrically and have been confused on Costa Rica's Osa
Peninsula. However, the leaves of C. grandifolia
usually have a more distinct submarginal vein, it has fewer stamens,
and the anthers are often 2--3 times longer than wide rather than
about as long as wide. In many details, e.g., fruit shape and color,
number and size of stamens, leaf coloration, C.
skutchii comes much closer to C. allenii.
Chrysochlamysskutchii is distinguished from C.
allenii by its larger, thinner leaves, glabrous inflorescence,
unequal outer sepal pair, persistent sepals in fruit, and lower
elevation habitat. The epithet is in honor of the species' first
collector, one of Costa Rica's primier natural historians, Alexander
Skutch.

Because of its apparently axillary inflorescences, bud-scale
scars, and resin-producing staminate flowers, this species's position
in Chrysochlamys is somewhat dubious. It has many close and
very prominent lateral veins, and small, essentially terminal
inflorescences that are often left in an apparently axillary position
(only one per node) by the dominant axillary vegetative shoot. It
also sometimes has unambiguously terminal inflorescences, and has an
androecium with the stamens more or less connivent into an apparently
resiniferous capitulum, much as in the resin-producing flowers of
some species of Clusia. The staminodia of the pistillate
flowers are connate into a ring or collar around the base of the
ovary as in Chrysochlamysmembranacea. The small,
narrowly ellipsoid-falcate, capsulate fruits mostly have only three
sessile stigmas, and usually one seed with a markedly cellular aril.
In leaf venation and bud scales it is very similar to
Symphonia, but for numerous reasons would be even more out of
place there than in Chrysochlamys. The species must certainly
belong to the Tovomita group, but further study is in order.
Although D'Arcy (1980) created the combination Tovomitopsismicrantha (Engler) D'Arcy for this material, the Peruvian
Chrysochlamysmicrantha Engler, by its original
description and type photo, bears it no resemblance.

Distribution. This species is known from scattered
localities in very wet forest on the Atlantic slope of Costa Rica and
Panama, and from a few collections from wet forest on the Burica
Peninsula, shared by both countries along their Pacific border.
Although most material is from below 800 m, the type is from nearly
1400 m.

The type and all other Costa Rican, Atlantic slope collections, as
well as one Panamanian collection from Bocas del Toro
(Kirkbride&Duke621), one from
Veraguas (McPherson10724), and two from the Río
Blanco de Norte area of the Atlantic lowlands of Coclé, have
particularly long, narrow leaves without obvious intersecondaries and
relatively few-flowered, very lax inflorescences with the outer
sepals slightly to very narrowly triangular. The Burica Peninsula
outliers and all other collections from Panama tend to have somewhat
broader and shorter leaves with more obvious intersecondaries,
multi-flowered, more congested inflorescences, and broader outer
sepals.

Chrysochlamystenuis is similar to C.
glauca in its dull, few-veined leaves and small flower buds.
The Río Guanche and Cerro Campana populations cited here were
discussed by D'Arcy (1980, p. 1034) under that species as "suggestive
of a distinct variety or even species." Rather than further blurring
what then seemed only suggestive, more recent collections,
particularly those from Atlantic Costa Rica, strengthen the case for
recognizing this entity as a distinct species. Other than the
narrower and usually reddish tan vs. gray-green-drying leaves, more
obscure venation and smaller, more delicate inflorescences, C.
tenuis is distinguished from C. glauca by the
resin canals of the leaves, which, when visible, are more or less
parallel to the midrib or at least somewhere crossing the lateral
veins, rather than parallel to the lateral veins, as in C.
glauca and most other species. The narrowly obovoid fruits of
C. tenuis are usually only 1- or 2-seeded, whereas the
more or less globose or broadly obovoid fruits of C.
glauca are normally 5-seeded. Morietal.
3950 recorded the seeds as "steel blue." The new species also
superficially resembles the Panamanian C. angustifolia,
which, however, is an epipetric rheophyte with milky resin, stouter
inflorescence axes, and much larger flower buds.

Original field notes for the type of this species indicated four
duplicates. These were earlier distributed as either
Tovomitopsisangustifolia or T. glauca,
and their whereabouts, except as indicated above, are uncertain.

Although the strictly South American Balboa does not occur
within the geographic focus of this paper, taxonomic clarity and
logical consistency prescribe its mention here as an additional
synonym of Chrysochlamys. Planchon and Triana described the
genus and its single species, B. membranaceae based
only on staminate material and placed it in their subtribe
"Euclusieae" rather than Tovomiteae. Examination of type material and
more recent collections, of both staminate and pistillate plants
(e.g., Boyle&Boyle3303 (MO),
Croat73004 (MO), Gentryetal.
14655 (MO), Hammel&Trainer15831 (MO), Hoover 1287 (MO), Hooveretal. 3136 (MO), Romero-Castañeda2739 (COL), Rubio&Quelal1339 (MO)) support the view that B. membranacea
is a species of Chrysochlamys:

This rare, somewhat problematic species might appear to walk the
generic borderline between Chrysochlamys and Tovomita.
The outer pair of sepals, although thinly membranaceous, exceed the
bud as in Tovomita, yet styles are lacking and the fruits
apparently do not have the dark maroon placenta and inner fruit wall
typical of Tovomita. Nevertheless, two Costa Rican collections
(Grayum3374, Grayum&Schatz5276) clearly indicate the presence of side shoots with
clustered leaves. With present knowledge, placement of this species
in Tovomita, particularly because of its branching pattern, is
more justifiable than placement in Chrysochlamys. As a
species, it is very distinctive, not only for the nearly sessile and
sometimes very large leaves, but also for the numerous (16--22)
lateral veins that are loop-connected to a distinct submarginal vein.
Probably because of the species' rarity, its short stature and large
leaves, the typical Tovomita branching pattern of T.
croatii has gone unobserved and unrecorded.

Acknowledgments. This work was supported, in part, by funds
from the National Science Foundation through a grant (DEB-9300814) to
the author and co-PI M. H. Grayum for the Manual to the Plants of
Costa Rica. I thank Mike Grayum for his always useful comments on an
early draft of the manuscript and Sylvia Troyo for the line drawings
of the two new species.